Detecting cryptic species in sympatry and allopatry: analysis of hidden diversity in Polyommatus (Agrodiaetus) butterflies (Lepidoptera: Lycaenidae)

Modern multilocus molecular techniques are a powerful tool in the detection and analysis of cryptic taxa. However, its shortcoming is that with allopatric populations it reveals phylogenetic lineages, not biological species. The increasing power of coalescent multilocus analysis leads to the situation in which nearly every geographically isolated or semi‐isolated population can be identified as a lineage and therefore raised to species rank. It leads to artificial taxonomic inflation and as a consequence creates an unnecessary burden on the conservation of biodiversity. To solve this problem, we suggest combining modern lineage delimitation techniques with the biological species concept. We discuss several explicit principles on how genetic markers can be used to detect cryptic entities that have properties of biological species (i.e. of actually or potentially reproductively isolated taxa). Using these principles we rearranged the taxonomy of the butterfly species close to Polyommatus (Agrodiaetus) ripartii. The subgenus Agrodiaetus is a model system in evolutionary research, but its taxonomy is poorly elaborated because, as a rule, most of its species are morphologically poorly differentiated. The taxon P. (A.) valiabadi has been supposed to be one of the few exceptions from this rule due to its accurately distinguishable wing pattern. We discovered that in fact traditionally recognized P. valiabadi is a triplet of cryptic species, strongly differentiated by their karyotypes and mitochondrial haplotypes.     

The genus Cyrtomium (Pteridophyta: Dryopteridaceae) in Africa and Madagascar

The fern genus Cyrtomium (Dryopteridaceae) in Africa and Madagascar is reviewed. Until now, a single Cyrtomium species ascribed to either C. caryotideum or C. micropterum (the latter often considered to be a variety of the former) was recognized for the region. This study shows that three Cyrtomium species occur in the region. One, C. falcatum, is not native to the African or Madagascan region, but has become naturalized in various parts of South Africa and Réunion. Two, formerly considered as a single taxon and confused with C. caryotideum and Indian forms thereof, are described as new – C. luctuosum from Madagascar, East, West and South Africa and C. pseudocaryotideum from South Africa. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167 , 449–465.     

The Biogeography of Cladocera (Crustacea) in Tropical Australia

Ninetysix species of Cladocera, comprising 8 Sididae, 50 Chydoridae, 17 Macrothricidae, 5 Moinidae, 2 Bosminidae, and 14 Daphniidae, occur in tropical Australia. Of this number 38% are cosmopolitan including so-called cosmopolitan species, 13.5% are circumtropical, 13.5% occur as well innear by countries, and 35% are endemic. North Queensland is a major area of speciation. Species richness varies from north to south and from wetter coastal areas to the drier inland. Major factors influencing richness and distribution are the relative proportion of floodplain billabongs in an area, coupled with rainfall reliability. The most common species include Alonella clathratula, Chydorus nr. pubsecens, Macrothrix triserialis, Oxyurella singalensis, Ephemeroporus nr. barroisi, Dadaya macrops, Diaphanosoma sarsi, Ceriodaphnia cornuta, and Dunhevedia crassa.     

Genome organization and polyploid evolution in the genus Eleusine (Poaceae)

 Eleusine (Poaceae) includes six diploid and three polyploid species and has three basic chromosome numbers, x=8, 9 and 10. The species are annual as well as perennial and all are wild except E. coracana, which is cultivated for grain and fodder in Africa and the Indian subcontinent. Eleusine coracana and E. africana have the same genome and chromosome number (2n=36). Eleusine indica and E. floccifolia are identified as two genome donors to these polyploid species. Eleusine kigeziensis is the third polyploid species of the genus with 2n=38. Its genome may have come from E. jaegeri and from one of the species with x=9, most probably from E. indica. Eleusine indica, E. tristachya, E. floccifolia and E. intermedia with x=9 and two polyploid species, E. coracana and E. africana, are closely related and there is free genetic flow between them. Eleusine multiflora with x=8 is significantly different in morphology and at genomic level from other species. Eleusine jaegeri with x=10 is morphologically similar to E. indica, however, more information is needed to ascertain its position in the genus. Eleusine coracana, which is commonly called finger millet, is a potential and nutritious crop for the increasing population of the world, particularly in arid and semi-arid regions. It can also serve as a gene pool for various important characters and disease resistant genes. Received February 11, 2002; accepted May 27, 2002 Published online: October 14, 2002 Addresses of the authors: Madho Singh Bisht and Yasuhiko Mukai (e-mail: ymukai@cc.osaka-kyoiku.ac.jp), Laboratory of Plant Molecular Genetics, Division of Natural Science, Osaka Kyoiku University, 4-698-1 Asahigaoka, Kashiwara, Osaka 582-8582, Japan.     

Invertebrate communities associated with a native (Vallisneria americana) and an alien (Trapa natans) macrophyte in a large river

1. We used a corer and a Downing box sampler to sample macroinvertebrates living on and beneath the introduced Trapa natans and the native Vallisneria americana in the freshwater tidal Hudson River, New York. 2. Densities of macroinvertebrates were higher in Trapa than in Vallisneria, and higher in the interior of plant beds than at their edges. These effects were largely a result of high plant biomass in Trapa beds and in bed interiors (the plants have similar surface area per unit mass). 3. The composition of both epiphytic and benthic macroinvertebrates differed distinctly between Trapa and Vallisneria, and also seasonally. 4. These compositional differences were not easily interpretable as rising from possible differences in oxygen concentrations, fish predation, or water circulation in the two macrophytes. 5. Sida crystallina (Cladocera) collected from Trapa contained more haemoglobin than those collected from Vallisneria. 6. The replacement of Vallisneria by Trapa in the Hudson probably increased system‐wide biodiversity and food for fish, although macroinvertebrates in Trapa beds may not be readily available to fish because of low oxygen concentration there.     

The Genus Rhyacophila Pictet (Trichoptera: Rhyacophilidae) in Vietnam

Currently, 20 species of the genus Rhyacophila are known from Vietnam. Of these, 17 species are distributed among eight species groups, with three species unassigned. Examination of material collected by personnel of the Royal Ontario Museum in Toronto, Canada has revealed six new species and five new records of Rhyacophila from Vietnam, distributed among six species groups, four of which are new to Vietnam. Newly described species include: Rhyacophila smithi, Rh. curriei, Rh. laocai, Rh. meyi, Rh. olahi, and Rh. fernandi. Rhyacophila bidens Kimmins, Rh. inaequalis Denning and Schmid, Rh. lhabu Schmid, Rh. malayana Banks, and Rh. rhombica Martynov are recorded from Vietnam for the first time. The females of four species, Rh. bidens, Rh. immaculata Mey, Rh. malayana Banks, and Rh. tamdaona Oláh, are described for the first time. All 31 species of Rhyacophila now known from Vietnam are included in this work, organized by branch and species group.     

Taxonomy of Electrogena antalyensis (Kazanci & Braasch, 1986) (Ephemeroptera,Heptageniidae)

Female imagines, subimagines, eggs and larvae of Electrogena antalyensis (Kazanci & Braasch) from Turkey are described for the first time. The results of a thorough analysis of larvae, based on the standard set of diagnostic characters for the identification of Electrogena species, are reported. The peculiarity of several characters places E. antalyensis in an isolated position within the genus Electrogena.     

The recent New Zealand species of Triforis (Gastropoda: Triforidae)

Paramendax apicina Powell and Mendax attenuatispira Powell are referred to genus Triforis Deshayes; T. antepallaxa, T. blacki, and T. tui are described as new; and a species close to the South Australian Triforis epallaxa (Verco) is recorded from New Zealand waters. Paramendax Powell is transferred from the Cerithiopsidae to the Triforidae as a subgenus of Triforis. It is suggested that the Recent South African Cerithiella nonnitens Barnard is referable to Triforis.     

Tracing the introduction history of the brown seaweed Dictyota cyanoloma (Phaeophyceae,Dictyotales) in Europe

Dictyota cyanoloma has recently been described from the Mediterranean Sea and Macaronesia but doubt had arisen as to whether this species was truly native in Europe. The species is mainly found on non-natural substrata (harbour walls, marinas, boat hulls, etc.), strongly suggesting that it is an introduction. Molecular sequence information from historical herbarium samples proves the presence of D. cyanoloma in the Adriatic Sea as early as 1935. Since approximately the year 2000, however, the number of records as well as the geographic range of the species has expanded significantly. The present-day distribution of D. cyanoloma occupies most of the Mediterranean Sea, Macaronesia, NW Africa and southern Portugal, but recent records from Galicia and SW England (Falmouth, Cornwall) indicate that the species is rapidly expanding northward. Collections from Australia demonstrated that the species is also present from Perth in Western Australia, over much of the southern Australian coastline up to Minnie Water in New South Wales. Phylogenetic analyses resolve D. cyanoloma in a sister clade to a previously unreported Australian Dictyota species. Analysis of genetic diversity of the mitochondrial markers (nad6–nad11 and atp9–orf11) reveals that even though Australian populations contain a much higher haplotype richness, European populations are also fairly diverse. Furthermore, only two out of 25 haplotypes are shared between both regions. These somewhat counterintuitive results could be indicative of a more complicated introduction history.     

Phylogeny and species reassessment of Hyalopterus (Aphididae,Aphidinae)

The broadly distributed genus Hyalopterus currently comprises three formally recognized species that are highly similar morphologically and hence difficult to be identified with certainty. This group has undergone multiple revisions in the past century, but none of these has assessed species from Asia, which has hampered our understanding of the species diversity within this genus. Based on a comprehensive data set from morphological data and host-associated data, and by coalescent-based delimitation approaches, the Hyalopterus species boundaries, distribution and diversity were clarified here to further reveal the composition of the species. Two single-locus (ML-GMYC and mPTP) and two multilocus (BPP and STACEY) delimitation methods were conducted based on extensive sampling. Then, the phylogenetic relationships and morphological divergence were assessed. Our data strongly supported that the number of recognized species in Hyalopterus had likely been underestimated. The phylogenetic analyses recovered four major clades, which corresponded to distinct host-plant preferences. Also, the morphological analyses showed significant differentiation for only one of the newly recognized candidate species uncovered by the delimitation approaches, suggesting the existence of at least two independent evolutionary lineages within Hyalopterus arundiniformis, which showed different patterns of host association. Moreover, based on our data, the taxonomic misidentification of H. arundiniformis in China was corrected here. This study lays the groundwork for the thorough taxonomic revision of Hyalopterus and for future evolutionary studies and underlines the importance of an integrated framework for species determination.     

Origins and implications of apid bees (Hymentopera: Apidae) in French Polynesia

Island plant–pollinator networks are typically simpler than their continental counterparts and this can make them less resilient to disturbance from exotic species. French Polynesia has a very low diversity of bees, but their status as either native or introduced species has been largely speculative. We combine previous studies with new DNA sequence data to show that 11 bee species have now been recorded for French Polynesia. Haplotype variation at the ‘barcode’ region of the mitochondrial gene cytochrome c oxidase subunit I (COI) for four of these species, Ceratina dentipes Freise, Xylocopa sonorina Smith, Braunsapis puangensis (Cockerell) and Amegilla pulchra (Smith), indicates that they all represent very recent introductions. Apis mellifera Linnaeus was a purposefully introduced species, and four megachilid species probably arrived due to human‐aided dispersal through maritime activities in the Pacific. The two remaining bee species, an unidentified partial specimen of a halictid bee and the colletid bee Hylaeus (P.) tuamotuensis Michener, are collectively known from only four specimens collected in the 1930s and their provenance is uncertain. French Polynesia therefore comprises a region where recently introduced bee species greatly overwhelm any possible native bee fauna. These introductions are likely to have major ecosystem impacts, including disruptions of existing plant–pollinator networks and facilitating the spread of weedy plant species, as well as positive impacts for agriculture. Future biosecurity initiatives need to consider these potential impacts and the likely routes of dispersal to effectively control any further unintended introductions.     

Invasion status of Florida bass Micropterus floridanus (Lesueur, 1822) in South Africa

Largemouth bass Micropterus salmoides are a popular North American angling species that was introduced into South Africa in 1928. To enhance the largemouth bass fisheries, Florida bass Micropterus floridanus were introduced into KwaZulu Natal, South Africa, in 1980. Knowledge on the status of M. floridanus in South Africa is required, because it lives longer and reaches larger sizes than M. salmoides, which may result in heightened impacts on native biota. Because M. floridanus are morphologically similar, but genetically distinct from M. salmoides, the distribution of this species was assessed by genetically screening 185 Micropterus sp. individuals sampled from 20 localities across South Africa using the mitochondrial ND2 gene. Individuals with mitochondrial DNA matching M. salmoides were recovered from 16 localities, whereas M. floridanus mitochondrial DNA was recovered from 13 localities. At nine localities (45%), the mitochondrial DNA of both species was detected. These results demonstrate M. floridanus dispersal to multiple sites across South Africa.     

Chromosomal Analysis of Neohydrocoptus jaechi (Wewalka) and Canthydrus diophthalmus (Reiche & Saulcy) (Coleoptera: Noteridae)

Neohydrocoptus jaechi (Wewalka) is recorded for the first time in Egypt. It is very similar to N. angolensis (Peschet), but there are aedeagal differences. The karyotype consists of 6 pairs of autosomes and sex chromosomes which are neo-XY (♂) and neo-XX (♀). C-banding shows that the sex chromosomes of Canthydrus diophthalmus (Reiche & Saulcy) are not the small pair suggested by Bilton (1992), but are one of the largest pairs, with the Y-chromosome totally heterochromatic. The implications of these findings are considered in the light of Belkaceme's (1991) work on the phylogeny of Noteridae. Neohydrocoptus is placed among the group of more primitive genera, outside the closely-knit group of higher Noteridae, while Canthydrus is placed right at the top of the higher Noteridae. The neo-XY sex chromosomes of Neohydrocoptus could be a primitive condition for the family, while the system found in Canthydrus is derivable from that of Synchortus Sharp (Saleh Ahmed et al., 1997) (one of Belkaceme's basal genera of the higher Noteridae) by loss of the mainly heterochromatic X₂-chromosome, leaving the more normal X₁-chromosome and the Y, which is almost entirely heterochromatic in Synchortus and completely so in Canthydrus.     

Genetic and chaetal variation in Nais worms (Annelida,Clitellata, Naididae)

The genus Nais is a group of oligochaetous clitellates, common in eutrophic freshwater habitats. About 30 species are described. Species identification is based primarily on chaetal characters, which are often subtle, inconsistent, and even overlapping between nominal species. We investigated the correlation between genetic variation and chaetal morphology in this genus. Eighty‐one individuals from Europe, North America, and China were included in the study. Seventy‐five of these were preserved as vouchers. They were scrutinized with regard to chaetal morphology, and ten different morphotypes were identified. Three molecular markers, two mitochondrial (the COI gene and 16S rDNA) and one nuclear (the ITS region), were used to establish the genetic lineages in the material. Genetic variation was found to be largely congruent with chaetal character patterns. However, at least nine separately evolving lineages (all supported by mitochondrial as well as nuclear data) correspond to at most six nominal species. Four morphotypes/lineages are recognized as Nais barbata, Nais christinae, Nais elinguis, and Nais stolci, respectively, whereas five, or possibly more, lineages represent a morphological continuum covering the variation of the Nais communis/variabilis complex. Thus, cryptic speciation is revealed. Our results indicate that a taxonomic revision of the genus will be needed in the future.     

Diversity of Siphonaria Sowerby I, 1823 (Gastropoda,Siphonariidae) in the Seychelles Bank and beyond

Molecular phylogenetic studies have shown that the characters of the reduced shell of the false limpets of the genus Siphonaria Sowerby I, 1823 are highly variable and often insufficient for species delimitation. The taxonomy and distribution of Siphonaria in the Indian Ocean are poorly known. We sampled Siphonaria in the Seychelles Bank to check the occurrence of recorded species using DNA sequences and to study the paths through which Siphonaria species have colonised the Seychelles Bank by reconstructing their phylogenetic relationships. Analyses of a dataset comprising 16 S rRNA gene sequences of 33 specimens from the Seychelles Bank and 300 additional Siphonaria sequences from other regions from GenBank with various methods for species delimitation resulted in 19–102 primary species hypotheses. Assemble Species by Automatic Partitioning provided a conservative estimate of the species number (42) in which several indisputable species were lumped. The results of Automatic Barcode Gap Discovery depended strongly on the assumed prior maximum intraspecific divergence, whereas the tree-based methods Generalised Mixed Yule Coalescent and Poisson Tree Processes resulted in high overestimates. The specimens from the Seychelles Bank represent three clades, belonging to the Siphonaria ‘atra’ group, the Siphonaria ‘normalis’ group and a possibly undescribed species recorded previously only from Hainan. At least two of the three species recorded from the Seychelles Bank came from the east, i.e., from the Coral Triangle in the Indo-Australian Archipelago, the region with the highest marine biodiversity worldwide. A major transport mechanism across the Indian Ocean was probably the South Equatorial Current.     

Four New Genera and Species of Chironomidae (Diptera) from Southern Africa

Four new genera are described for four new species of Chironomidae from southern Africa. Paradoxocladius mangoldi (Orthocladiinae) is based on all life stages, Lunditendipes tropicum (Chironominae) on male adults, Xestotendipes sufflavus (Chironominae) on male and female adults, and Zuluchironomus demoori (Chironominae) on male and female adults. A highly apomorphic larva with a cleft mid-mentum, possibly belonging to Z. demoori, is also described. Some ecological details are given.     

Discovery of the tribe Odontalgini (Coleoptera: Staphylinidae; Pselaphinae) in Japan

A new species of the pselaphine genus Odontalgus is described from Honshu, Japan, under the name of Odontalgus masaoi. Odontalgus coreanus Löbl known from North Korea is recorded for the first time from Kyushu, Japan. These are the first formal records of the tribe Odontalgini from Japan.     

Predation of Thereva nobilitata (Fabricius) (Diptera: Therevidae) on Agriotes obscurus L. (Coleoptera: Elateridae)

Little is known about the natural enemies of wireworms (Coleoptera: Elateridae), but there are frequent anecdotal reports of (usually unnamed) stiletto fly larvae (Diptera: Therevidae) preying on various species. We observed larvae of Thereva nobilitata (Fabricius) feeding on larvae of the dusky wireworm, Agriotes obscurus L., during the summer of 2011, in Agassiz, British Columbia. This finding is of interest as: both the predator and the wireworm are introduced species to this area from Europe; T. nobilitata is uncommon in North America; and this predator has not been associated with any wireworm species previously. We observed that larvae of male and female T. nobilitata will feed on various sizes of A. obscurus larvae, most feeding being carried out by the smallest T. nobilitata larvae. These findings suggest future work should assess the potential for therevid larvae as top‐down regulators of Agriotes larvae under field conditions.     

Notes on the genus Agnetina (= Phasganophora) (Plecoptera: Perlidae)

Genus Agnetina Klapálek, 1907 is removed from synonymy with Dinocras Klapálek, 1907 and synonymy between Agnetina and Phasganophora Klapálek, 1914 is established. A list of presently recognised nominal species of Agnetina is provided. For all west palaearctic species, i.e., A. elegantula (Klapálek), A. senilis Klapálek, A. werneri (Kempny), comb, n., lectotypes are designated. Distinctive characters of these three species are described, external male genitalia and eggs are illustrated. Asian A. brevipennis (Navás), comb, n., is briefly compared. A. dubia nom. n. is proposed to replace A. brevipennis Klapálek, 1921 (not Navás, 1912), a doubtful Asian species. A. pedata (Koponen, 1949) and A. undaata (Klapálek, 1921) are considered possible synonyms of A. senilis. which is for the first time recorded from Central Asia (Baikal area).